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Restrepo et al ANNOTATED LIST OF SPECIES Check List 13 (4): 203–237 https://doi.org/10.15560/13.4.203

Amphibians and reptiles from two localities in the northern Andes of Colombia

Adriana Restrepo, Claudia Molina-Zuluaga, Juan P. Hurtado, Carlos M. Marín, Juan M. Daza

Grupo Herpetológico de Antioquia, Instituto de Biología, Universidad de Antioquia, A.A. 1226, Medellín, Colombia. Corresponding author: Juan M. Daza, [email protected]

Abstract The northern Andes are a geographically, climatically, and ecologically complex area that harbors a high number of species and presents a high level of endemism. However, the landscape of this region has been highly transformed by human activities, including agriculture, cattle grazing, urbanization, and recently, the construction of large infrastruc- ture projects. This study presents a list of amphibians and reptiles compiled from field surveys carried out between 2006 and 2016 in the protected areas surrounding 2 hydroelectric projects located on the eastern flank of the northern Cordillera Central in Colombia. The checklist is comprised of 101 species (43 amphibians and 58 reptiles), including 17 endemic to Colombia and 4 threatened species. The high species density, high representativeness of the regional biodiversity, and high level of amphibian endemism make these 2 protected areas important for conservation of the regional herpetofauna.

Key words Endemism; frogs; hydroelectric dam; lizards; snakes; species richness.

Academic editor: Sebastian Lotzkat | Received 3 December 2015 | Accepted 4 June 2017 | Published 28 July 2017

Citation: Restrepo A, Molina-Zuluaga C, Hurtado JP, Marín CM, Daza JM (2017) Amphibians and reptiles from two localities in the northern Andes of Colombia. Check List 13 (4): 203–237. https://doi.org/10.15560/13.4.203

Introduction the final closure of the Isthmus of Panama that allowed the biotic interchange between 2 continents, and (3) the The Andes mountain range extends for 5000 km along climatic fluctuations during the Pleistocene that lead to the western coast of South America and 3 main domains cycles of range contraction and expansion resulting in the are recognized: southern, central and northern Andes fragmentation and isolation of natural populations, with (Gregory-Wodzicki 2000). The northern Andes reach subsequent speciation and radiation events (Duellman their maximum geomorphological complexity in Colom- 1999, Kattan et al. 2004). Thus, the diversity of ecologi- bia, where 3 mountain chains (the Western, Central, and cal settings generated by this physiographic complexity Eastern cordilleras), are separated by the sedimentary has made the northern Andes one of the most biodiverse basins of the Cauca and Magdalena rivers (Kattan et al. regions on the planet with an exceptional number of 2004). Three major historical events have shaped the bio- endemic plants and vertebrates (Fjeldsa 1994, Duellman logical diversity in this region: (1) the uplift of mountain 1999, Myers et al. 2000). ranges in a complex series of orogenic processes generat- The northern Andes are also under continuous and ing new environments and causing vicariant events, (2) intense anthropogenic pressure. About 70% of the human

Copyright Restrepo et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 204 Check List 13 (4)

Figures 1. Study area showing the location in Colombia and the northern Cordillera Central. On the right, the two localities (Jaguas and San Carlos) are expanded and black dots represent the sampling sites. Colors represent cover type. population in Colombia is concentrated in this region and Methods agricultural activities, cattle grazing and infrastructure development (i.e. urbanization, dam construction, and Study sites. This study was conducted in the northern roads) are among the main causes of habitat loss and Andes at 2 sites above 500 m elevation in the Cordil- fragmentation (Etter and van Wyngaarden 2000, Fjeldsa lera Central in Antioquia department, Colombia (Fig. et al. 2005). On the eastern flank of the northern Cordil- 1). The Jaguas site (06°26ʹ06.0ʺ N, 075°00ʹ57.6ʺ W) lera Central, several hydroelectric projects involving dam includes a 4000 ha private protected area with habitats in constructions, roads, and power lines were developed different successional stages, within the premontane wet in the last 2 decades. As a mitigation strategy, Colom- forest life zone (bmh-PM, Holdridge 1964). The altitude bia’s environmental legislation demands that these large ranges between 1100 and 1300 m above sea level. The temperature ranges between 18 and 24 °C and the annual infrastructure projects compensate the negative impacts rainfall between 2000 and 4000 mm. The San Carlos through the implementation of effective conservation and site (06°12ʹ57.6ʺ N, 074°51ʹ21.6ʺ W) includes a private restoration plans by preserving and monitoring the biodi- protected area of 3000 ha, within the tropical moist forest versity in the surrounding areas (Minambiente 2012). As life zone (bh-T, Holdridge 1964). The altitude is between a consequence, these habitats, as virtually the only rem- 600 and 950 m above sea level. It has a mean temperature nants left, represent a last-minute opportunity to make of 24 °C and the annual rainfall ranges between 2000 and complete species checklists for the northern portion of 4000 mm. The 2 sites are under a bimodal rainfall regime the central Andes of Colombia. with 1 rainy season between March and May and 1 Based on more than a decade of field surveys, we pro- between September and November (Poveda et al. 2005). vide the species list of amphibians and reptiles found in the protected areas of 2 main hydroelectric projects located on Data collection. Between 2006 and 2016, the complete the northeastern Cordillera Central in Colombia. species inventory has been an ongoing goal as part of the Restrepo et al. | Herpetofauna of the northern Andes of Colombia 205 monitoring plan implemented by the company ISAGEN (2000) proposed R. pseudopalmatus as sister species of S.A. Amphibians and reptiles were searched for in all R. palmatus. However, Grant et al. (2006) question the possible microhabitats including diurnal and nocturnal validity of morphological characters used to distinguish sampling events during rainy, dry, and transition seasons. this species from R. palmatus and consider that both spe- In total, 49 sampling points were distributed in different cies could be conespecific given that the type locality of vegetation types. R. pseudopalmatus (Amalfi, Antioquia) lies within the Collected specimens were euthanized with 2% Roxi­ known distribution of R. palmatus on the eastern slope cain®, fixed in 10% formalin solution, and preserved of the Central Cordillera. Here, we assign our samples in 70% ethanol following the protocol by Cortez et al. to R. pseudopalmatus according to Muñoz-Ortíz et al. (2006). Voucher specimens are deposited in the Museo (2015) who suggested, based on molecular information, de Herpetología Universidad de Antioquia (MHUA), in that this species could be redescribed restricting R. pseu- Medellín, Colombia (Instituto Alexander von Humboldt dopalmatus to the Central Cordillera and R. palmatus to registry # 80, since the year 2000). All procedures were the Eastern Cordillera. conducted under permit No. 112–0046 granted by the Rhinella alata (Thominot, 1884): Figure 2B Corporación Autónoma Regional CORNARE. We list the Bufo alatus Thominot 1884: 151. collected and examined specimens in Table 1 (amphib- Rhinella alata — Frost et al. 2006: 365 ians) and 2 (reptiles). All coordinates use the WGS84 Material examined. Table 1. datum. Rhinella alata is a small-sized toad with skin of Amphibian nomenclature follows Frost (2016) and dorsum bearing a mixture of warts, pustules and minute reptile nomenclature follows Uetz and Hošek (2016). We tubercles; supraorbital crests low and thick, continuous reviewed the original descriptions of the species along with preorbital crests; dorsal coloration ranging from with the MHUA reference collection to corroborate the light gray or brown to dark gray or brown, with irregular taxonomic identifications. We included in the list only black and yellowish marks (a mid-dorsal line is usu- the species having voucher specimens or photographs ally present) and coloration of ventral surfaces cream to that allow unambiguous identification of taxa. We follow yellowish-cream with irregular darker marks arranged in the species distribution provided by IUCN (2016) for diverse patterns (dos Santos et al. 2015). Populations of amphibians and Uetz and Hošek (2016) for reptiles. Rhinella margaritifera species group distributed on Mag- dalena river valley have commonly been referred to as Results R. margaritifera (see Burbano et al. 2016). Nevertheless, although R. alata is closely related to R. margaritifera Amphibians (distributed in the upper Amazon basin of Peru and From the 2 protected areas, we report 43 amphibian spe- Ecuador), this species may be easily distinguished by dif- cies representing 13 families and 25 genera (Table 3). ferences in body size (R. margaritifera being significant In the protected area of the Jaguas locality, we report 37 larger than R. alata) and the relative size of the cranial amphibian species from 11 families. The most diverse crest (larger in R. margaritifera than in R. alata). lineages were Hylidae with 12 species (32%) and Crau- Rhinella horribilis (Wiegmann, 1833): Figure 2C gastoridae with 6 species (16%). Eleven species are Bufo horribilis Wiegmann 1833: 654. endemic to Colombia (30%), and according to the IUCN Bufo marinus — Peters 1873: 618. Red Lists, 1 is classified as Endangered, 2 as Vulnerable, Rhinella horribilis — Acevedo et al. 2016: 584. and 1 as Near Threatened (IUCN 2016). In comparison, Material examined. Table 1. the San Carlos site has 34 amphibian species grouped in Recently, Acevedo et al. (2016) proposed that popu- 11 families. Again, the most diverse groups were Hylidae lations of Rhinella marina (Linnaeus, 1758) distributed with 11 species (32%) and Craugastoridae with 6 spe- west of the Andean cordilleras should be considered a cies (18%). Eleven species from San Carlos locality are different species from the eastern populations. There- endemic to Colombia (32%) and 2 are listed as Vulnerable fore, the eastern populations should maintain the name (IUCN 2016). Jaguas and San Carlos localities shared 28 R. marina and R. horribilis should be revalidated for amphibian species. the western populations. According to this concept, the Magdalena river valley populations correspond to R. hor- Rheobates pseudopalmatus (Rivero & Serna, 2000): ribilis (Wiegmann, 1833). Figure 2A Colostethus pseudopalmatus Rivero and Serna 2000: 55. Cochranella resplendens (Lynch & Duellman, 1973): Rheobates pseudopalmatus — Grant et al. 2006: 159. Figure 2D Material examined. Table 1. Centrolenella resplendens Lynch and Duellman 1973: 51. A moderate-sized frog with cryptic, brown or gray Cochranella resplendens — Ruíz-Carranza and Lynch 1991: 23. dorsal coloration; dorsal skin granular posteriorly; toe Material examined. Table 1. webbing extensive; finger I shorter than finger II; pale Espadarana prosoblepon (Boettger, 1892): Figure 2E oblique lateral stripe present or absent and pale dorso- Hyla prosoblepon Boettger 1892: 45. lateral and ventrolateral stripes absent. Rivero and Serna Centrolene prosoblepon Noble 1924: 66. 206 Check List 13 (4)

Table 1. Voucher list of amphibians of Jaguas and San Carlos hydroelectric protected areas with geographic coordinates.

Taxon Voucher Latitude Longitude Taxon Voucher Latitude Longitude AMPHIBIA MHUA-A 10688 6.19176 -74.80257 ANURA Hyalinobatrachium MHUA-A 07075 6.23108 -74.87661 Aromobatidae aureoguttatum MHUA-A 07076 6.23108 -74.87661 Rheobates pseudopalmatus MHUA-A 07845 6.38871 -75.02555 Hyalinobatrachium MHUA-A 07198 6.21734 -74.81262 MHUA-A 08485 6.36725 -75.02723 fleischmanni MHUA-A 08491 6.39128 -75.03640 Rulyrana susatamai MHUA-A 09539 6.36725 -75.02723 MHUA-A 09425 6.39128 -75.03640 Sachatamia punctulata MHUA-A 09728 6.32866 -75.01155 MHUA-A 09426 6.36725 -75.02723 MHUA-A 09764 6.29817 -74.91981 MHUA-A 09538 6.39128 -75.03640 MHUA-A 10803 6.28806 -74.82722 MHUA-A 09758 6.29817 -74.91981 MHUA-A 10804 6.28806 -74.82722 MHUA-A 10599 6.15750 -75.04325 Sachatamia sp. MHUA-A 07123 6.37141 -75.04253 MHUA-A 10641 6.15750 -75.04325 MHUA-A 07842 6.38871 -75.02555 MHUA-A 10643 6.15829 -75.04531 MHUA-A 07844 6.38871 -75.02555 MHUA-A 10644 6.15829 -75.04531 MHUA-A 08205 6.39202 -75.03650 MHUA-A 10646 6.15829 -75.04531 MHUA-A 08216 6.38539 -75.02202 MHUA-A 10650 6.15829 -75.04531 MHUA-A 08217 6.39170 -75.03626 MHUA-A 10651 6.15750 -75.04325 MHUA-A 08229 6.39202 -75.03650 MHUA-A 10652 6.15763 -75.04316 MHUA-A 08231 6.39202 -75.03650 MHUA-A 10653 6.15763 -75.04316 MHUA-A 08232 6.39202 -75.03650 MHUA-A 10654 6.15829 -75.04531 MHUA-A 08233 6.39202 -75.03650 MHUA-A 10662 6.15829 -75.04531 MHUA-A 08336 6.39115 -75.03641 MHUA-A 10726 6.20908 -74.85436 MHUA-A 08337 6.39115 -75.03641 Bufonidae MHUA-A 10615 6.37172 -75.04534 Rhinella alata MHUA-A 09731 6.32711 -75.01186 MHUA-A 10682 6.36662 -75.02798 MHUA-A 09732 6.32758 -75.01176 Craugastoridae MHUA-A 09739 6.27667 -74.93102 Craugastor metriosistus MHUA-A 10468 6.24144 -74.86976 MHUA-A 09766 6.29231 -75.03969 MHUA-A 10507 6.23974 -74.86880 MHUA-A 08492 6.39128 -75.03640 MHUA-A 10586 6.15763 -75.04316 MHUA-A 06979 6.30032 -75.06698 MHUA-A 10587 6.36767 -75.02635 MHUA-A 07930 6.38871 -75.02555 MHUA-A 10588 6.38518 -75.02140 MHUA-A 08225 6.36725 -75.02723 MHUA-A 10589 6.24144 -74.86976 MHUA-A 08330 6.36725 -75.02723 MHUA-A 10590 6.20938 -74.85352 MHUA-A 08486 6.39128 -75.03640 MHUA-A 10656 6.20908 -74.85436 MHUA-A 10671 6.18324 -74.79362 MHUA-A 10660 6.20908 -74.85436 MHUA-A 10673 6.18325 -74.79362 MHUA-A 10800 6.28806 -74.82722 MHUA-A 10674 6.18280 -74.19330 MHUA-A 10808 6.15844 -75.04433 MHUA-A 10676 6.19224 -74.80183 MHUA-A 10809 6.15865 -75.04447 MHUA-A 10677 6.15671 -75.04108 Pristimantis fallax MHUA-A 07125 6.20671 -74.85500 MHUA-A 10679 6.18336 -74.79308 MHUA-A 10523 6.20908 -74.85436 Rhinella horribilis MHUA-A 10722 6.13321 -75.09676 Pristimantis gaigei MHUA-A 07118 6.24320 -74.87138 Centrolenidae MHUA-A 07208 6.21854 -74.83727 Cochranella resplendens MHUA-A 9540 6.36725 -75.02723 MHUA-A 07209 6.21626 -74.83435 Espadarana prosoblepon MHUA-A 06978 6.30032 -75.06698 MHUA-A 08348 6.19062 -74.80474 MHUA-A 07124 6.36699 -75.02892 MHUA-A 08497 6.36725 -75.02723 MHUA-A 07420 6.36930 -75.02520 MHUA-A 09750 6.27681 -74.93975 MHUA-A 07421 6.36930 -75.02520 MHUA-A 09751 6.27679 -74.93085 MHUA-A 08212 6.38871 -75.02335 MHUA-A 10495 6.17844 -74.80273 MHUA-A 08213 6.38871 -75.02335 MHUA-A 10516 6.19224 -74.80183 MHUA-A 08214 6.38871 -75.02335 MHUA-A 10802 6.28806 -74.82722 MHUA-A 08226 6.36725 -75.02723 Pristimantis jaguensis MHUA-A 07026 6.38871 -75.02335 MHUA-A 08230 6.39202 -75.03650 MHUA-A 07027 6.38871 -75.02335 MHUA-A 08329 6.36725 -75.02723 MHUA-A 07028 6.38871 -75.02335 MHUA-A 08338 6.39115 -75.03641 MHUA-A 07030 6.38871 -75.02335 MHUA-A 08493 6.39128 -75.03640 MHUA-A 07031 6.38871 -75.02335 MHUA-A 09723 6.32845 -75.01171 MHUA-A 07032 6.38871 -75.02335 MHUA-A 09724 6.32734 -75.01160 MHUA-A 07033 6.38871 -75.02335 MHUA-A 09729 6.32734 -75.01160 MHUA-A 07238 6.38871 -75.02335 MHUA-A 09730 6.32734 -75.01160 MHUA-A 07246 6.38871 -75.02335 MHUA-A 10629 6.38518 -75.02140 MHUA-A 07248 6.38871 -75.02335 MHUA-A 10631 6.37172 -75.04534 MHUA-A 07249 6.38871 -75.02335 MHUA-A 10632 6.35150 -74.99528 MHUA-A 07250 6.38871 -75.02335 MHUA-A 10633 6.35150 -74.99528 MHUA-A 07251 6.38871 -75.02335 MHUA-A 10634 6.35150 -74.99528 MHUA-A 07252 6.38871 -75.02335 MHUA-A 10635 6.19176 -74.80257 MHUA-A 07253 6.38871 -75.02335 MHUA-A 10636 6.38518 -75.02140 MHUA-A 07254 6.38871 -75.02335 MHUA-A 10638 6.35150 -74.99528 MHUA-A 07255 6.38871 -75.02335 MHUA-A 10681 6.36662 -75.02798 MHUA-A 07256 6.35812 -74.99367 MHUA-A 10686 6.35150 -74.99528 MHUA-A 07257 6.35812 -74.99367 Continued Restrepo et al. | Herpetofauna of the northern Andes of Colombia 207

Table 1. Continued.

Taxon Voucher Latitude Longitude Taxon Voucher Latitude Longitude MHUA-A 07415 6.38871 -75.02335 MHUA-A 10503 6.15801 -75.04480 MHUA-A 07417 6.38871 -75.02335 MHUA-A 10522 6.15829 -75.04379 MHUA-A 07837 6.38871 -75.02555 MHUA-A 10525 6.15829 -75.04379 MHUA-A 10518 6.36662 -75.02798 MHUA-A 10591 6.24279 -74.86818 Pristimantis penelopus MHUA-A 07034 6.34818 -75.00237 Pristimantis viejas MHUA-A 07058 6.34818 -75.00237 MHUA-A 07035 6.34818 -75.00237 MHUA-A 07135 6.24739 -74.86938 MHUA-A 07424 6.36930 -75.02520 MHUA-A 08335 6.39202 -75.03650 MHUA-A 08588 6.36728 -75.02697 MHUA-A 08341 6.38528 -75.02158 MHUA-A 09727 6.32860 -75.01146 MHUA-A 08538 6.40999 -75.14157 MHUA-A 09733 6.32831 -75.01188 MHUA-A 09742 6.29610 -74.91944 MHUA-A 09734 6.32866 -75.01155 MHUA-A 09743 6.27721 -74.92910 MHUA-A 09754 6.29647 -74.91934 MHUA-A 09744 6.27721 -74.92910 MHUA-A 09756 6.29771 -74.91969 MHUA-A 09755 6.29631 -74.91931 MHUA-A 09760 6.29817 -74.91981 MHUA-A 09759 6.29648 -74.91931 MHUA-A 10461 6.15750 -75.04325 MHUA-A 10465 6.38518 -75.02140 MHUA-A 10462 6.36697 -75.02733 MHUA-A 10474 6.37198 -75.04549 MHUA-A 10463 6.36701 -75.02718 MHUA-A 10475 6.15750 -75.04325 MHUA-A 10464 6.37194 -75.04556 MHUA-A 10521 6.37198 -75.04549 MHUA-A 10466 6.36662 -75.02798 Dendrobatidae MHUA-A 10467 6.15728 -75.42740 Colostethus inguinalis MHUA-A 00223 6.21591 -74.85079 MHUA-A 10469 6.35150 -74.99528 MHUA-A 07072 6.22342 -74.84733 MHUA-A 10470 6.35150 -74.99528 MHUA-A 10645 6.18324 -74.79362 MHUA-A 10476 6.35150 -74.99528 MHUA-A 10600 6.18370 -74.79353 MHUA-A 10478 6.36662 -75.02798 MHUA-A 10642 6.18370 -74.79353 MHUA-A 10479 6.37188 -75.04533 MHUA-A 10647 6.19976 -74.80432 MHUA-A 10480 6.35150 -74.99528 MHUA-A 10648 6.18324 -74.79362 MHUA-A 10482 6.36697 -75.02733 MHUA-A 10649 6.24307 -74.86771 MHUA-A 10483 6.38518 -75.02140 MHUA-A 10661 6.18370 -74.79353 MHUA-A 10485 6.15822 -75.04457 Colostethus aff. fraterdanieli MHUA-A 07396 6.38871 -75.02335 MHUA-A 10486 6.15723 -75.04282 MHUA-A 07411 6.38871 -75.02335 MHUA-A 10487 6.36701 -75.02718 MHUA-A 07416 6.36930 -75.02520 MHUA-A 10488 6.37172 -75.04534 MHUA-A 07419 6.38871 -75.02335 MHUA-A 10490 6.15754 -75.04295 MHUA-A 08472 6.38528 -75.02158 MHUA-A 10491 6.36662 -75.02798 MHUA-A 08587 6.36728 -75.02697 MHUA-A 10494 6.37188 -75.04533 MHUA-A 09533 6.36725 -75.02723 MHUA-A 10496 6.15829 -75.04531 MHUA-A 09534 6.36725 -75.02723 MHUA-A 10497 6.15773 -75.04398 MHUA-A 09535 6.36725 -75.02723 MHUA-A 10500 6.37158 -75.04247 MHUA-A 09536 6.38537 -75.02166 MHUA-A 10501 6.38518 -75.02140 Dendrobates truncatus MHUA-A 08223 6.19055 -74.80480 MHUA-A 10502 6.36662 -75.02798 MHUA-A 07410 6.22020 -74.84713 MHUA-A 10504 6.20908 -74.85436 MHUA-A 08210 6.19570 -74.81670 MHUA-A 10505 6.20908 -74.85436 MHUA-A 09740 6.27738 -74.92780 MHUA-A 10506 6.15849 -75.04496 MHUA-A 10695 6.18360 -74.79343 MHUA-A 10510 6.15750 -75.04325 Eleutherodactylidae MHUA-A 10511 6.15829 -75.04531 Diasporus anthrax MHUA-A 07237 6.34818 -75.00237 MHUA-A 10513 6.35150 -74.99528 Diasporus gularis MHUA-A 07258 6.38871 -75.02335 MHUA-A 10515 6.15723 -75.04282 MHUA-A 07405 6.38871 -75.02335 MHUA-A 10517 6.15784 -75.04414 MHUA-A 07406 6.38871 -75.02335 MHUA-A 10519 6.37188 -75.04533 MHUA-A 07407 6.38871 -75.02335 MHUA-A 10520 6.36662 -75.02798 MHUA-A 07412 6.38871 -75.02335 MHUA-A 10527 6.36697 -75.02733 MHUA-A 07413 6.38871 -75.02335 MHUA-A 10559 6.15829 -75.04531 MHUA-A 07839 6.38871 -75.02555 MHUA-A 10560 6.37194 -75.04556 MHUA-A 08211 6.38871 -75.02335 MHUA-A 10592 6.20908 -74.85436 MHUA-A 08494 6.38528 -75.02158 MHUA-A 10724 6.20908 -74.85436 MHUA-A 08496 6.39128 -75.03640 MHUA-A 10725 6.20908 -74.85436 MHUA-A 10593 6.38518 -75.02140 MHUA-A 10811 6.13846 -75.06631 MHUA-A 10594 6.20908 -74.85436 Pristimantis taeniatus MHUA-A 07211 6.18737 -74.80460 MHUA-A 10595 6.20908 -74.85436 MHUA-A 07212 6.19743 -74.80363 MHUA-A 10794 6.37341 -75.02509 MHUA-A 07215 6.21569 -74.81371 Hylidae MHUA-A 07234 6.35447 -74.98836 Boana boans MHUA-A 07193 6.22895 -74.87296 MHUA-A 07423 6.36930 -75.02520 MHUA-A 07838 6.38871 -75.02555 MHUA-A 08484 6.36725 -75.02723 MHUA-A 07840 6.36664 -75.02746 MHUA-A 09537 6.37540 -74.98122 MHUA-A 09505 6.36081 -74.98657 MHUA-A 09757 6.29681 -74.91933 MHUA-A 09752 6.28320 -74.92413 MHUA-A 09769 6.29371 -75.04156 MHUA-A 10564 6.19175 -74.80257 MHUA-A 09770 6.29371 -75.04156 Boana xerophylla MHUA-A 10567 6.19725 -74.80433 MHUA-A 10471 6.15829 -75.04379 MHUA-A 10566 6.19725 -74.80433 Continued 208 Check List 13 (4)

Table 1. Continued.

Taxon Voucher Latitude Longitude Taxon Voucher Latitude Longitude MHUA-A 10568 6.19725 -74.80433 MHUA-A 10556 6.19725 -74.80433 MHUA-A 10569 6.19725 -74.80433 MHUA-A 10581 6.19725 -74.80433 MHUA-A 10571 6.18406 -74.79338 MHUA-A 10582 6.19725 -74.80433 Dendropsophus ebraccatus MHUA-A 07196 6.19744 -74.80397 MHUA-A 10583 6.19725 -74.80433 MHUA-A 10692 6.19725 -74.80433 MHUA-A 10584 6.19725 -74.80433 MHUA-A 10528 6.19725 -74.80433 MHUA-A 10585 6.15781 -75.04087 MHUA-A 10529 6.35171 -74.99379 Scinax sp. MHUA-A 07843 6.38871 -75.02555 MHUA-A 10532 6.35171 -74.99379 MHUA-A 08228 6.36081 -74.98657 MHUA-A 10533 6.35171 -74.99379 Scinax rostratus MHUA-A 07233 6.35812 -74.99367 MHUA-A 10535 6.19725 -74.80433 MHUA-A 10799 6.28806 -74.82722 MHUA-A 10538 6.19725 -74.80433 Scinax ruber MHUA-A 01685 6.49061 -74.83683 MHUA-A 10542 6.19725 -74.80433 MHUA-A 01686 6.49061 -74.83683 MHUA-A 10544 6.19725 -74.80433 MHUA-A 09500 6.36081 -74.98657 MHUA-A 10551 6.19725 -74.80433 Smilisca phaeota MHUA-A 08539 6.40999 -75.14157 MHUA-A 10553 6.19725 -74.80433 MHUA-A 09503 6.36081 -74.98657 MHUA-A 10555 6.35171 -74.99379 MHUA-A 09746 6.27700 -74.92966 MHUA-A 10557 6.35171 -74.99379 MHUA-A 09761 6.29143 -74.93671 Dendropsophus norandinus MHUA-A 08235 6.39202 -75.03650 MHUA-A 09762 6.29143 -74.93671 MHUA-A 08488 6.39128 -75.03640 MHUA-A 09767 6.29243 -75.04211 MHUA-A 08589 6.39214 -75.03689 MHUA-A 10572 6.15743 -75.04177 MHUA-A 08590 6.39214 -75.03689 MHUA-A 10573 6.38518 -75.02140 MHUA-A 08591 6.39214 -75.03689 MHUA-A 10574 6.24045 -74.86977 MHUA-A 10530 6.15784 -75.04414 MHUA-A 10575 6.24045 -74.86977 MHUA-A 10531 6.15784 -75.04414 MHUA-A 10576 6.35171 -74.99379 MHUA-A 10534 6.15750 -75.04325 MHUA-A 10577 6.24045 -74.86977 MHUA-A 10536 6.15801 -75.04480 MHUA-A 10578 6.35171 -74.99379 MHUA-A 10537 6.15784 -75.04414 MHUA-A 10579 6.35171 -74.99379 MHUA-A 10539 6.36798 -74.99639 MHUA-A 10580 6.15850 -75.04436 MHUA-A 10540 6.36798 -74.99639 MHUA-A 10727 6.20908 -74.85436 MHUA-A 10541 6.15750 -75.04325 Leptodactylidae MHUA-A 10543 6.15850 -75.04436 Engystomops pustulosus MHUA-A 09726 6.32913 -75.01398 MHUA-A 10545 6.15829 -75.04531 MHUA-A 10602 6.18406 -74.79338 MHUA-A 10546 6.15742 -75.04264 MHUA-A 10603 6.19725 -74.80433 MHUA-A 10547 6.15750 -75.04325 MHUA-A 10604 6.19725 -74.80433 MHUA-A 10548 6.15754 -75.04295 MHUA-A 10605 6.19725 -74.80433 MHUA-A 10549 6.36798 -74.99639 MHUA-A 10606 6.19725 -74.80433 MHUA-A 10550 6.36798 -74.99639 Leptodactylus colombiensis MHUA-A 07197 6.19144 -74.80397 MHUA-A 10554 6.36798 -74.99639 MHUA-A 09501 6.39128 -75.03640 MHUA-A 10561 6.15850 -75.04436 MHUA-A 10806 6.15651 -75.04146 MHUA-A 10562 6.15784 -75.04414 MHUA-A 10807 6.15651 -75.04146 Dendropsophus subocularis MHUA-A 07408 6.38871 -75.02335 Leptodactylus fuscus MHUA-A 01687 6.49061 -74.83683 MHUA-A 07409 6.38871 -75.02335 MHUA-A 10598 6.38747 -74.99274 MHUA-A 07834 6.38871 -75.02555 MHUA-A 10790 6.38788 -74.99348 MHUA-A 08206 6.39202 -75.03650 MHUA-A 10791 6.38788 -74.99348 MHUA-A 08339 6.39128 -75.03640 CAUDATA MHUA-A 08340 6.39128 -75.03640 Plethodontidae MHUA-A 08490 6.39128 -75.03640 Bolitoglossa ramosi MHUA-A 07899 6.39100 -75.02100 Hyloscirtus palmeri MHUA-A 06977 6.30032 -75.06698 GYMNOPHIONA MHUA-A 07073 6.23108 -74.84733 Caeciliidae MHUA-A 09504 6.36081 -74.98657 Caecilia thompsoni MHUA-A 07192 6.62890 -75.66244 MHUA-A 10563 6.15766 -75.04323 MHUA-A 07115 6.51013 -75.04172 MHUA-A 10693 6.23974 -74.86880 Siphonopidae MHUA-A 10805 6.15665 -75.04128 Microcaecilia pricei MHUA-A 10728 6.29648 -74.82229 MHUA-A 10810 6.15865 -75.04447 Rhinatrematidae Phyllomedusa venusta MHUA-A 07121 6.21284 -74.82631 Epicrionops parkeri MHUA-A 07191 6.22895 -74.87296 MHUA-A 07195 6.22895 -74.87296 MHUA-A 10720 6.20938 -74.85352

Espadarana prosoblepon — Guayasamin et al. 2009: 33. Small glass frog which exhibits complete ventral Material examined. Table 1. transparency; transparent pericardium; white hepatic and visceral peritonea; white bones; iris silvery at the Hyalinobatrachium aureoguttatum (Barrera-Rodríguez center, yellow at the edges with numerous black spots & Ruíz-Carranza, 1989): Figure 2F Centrolenella aureoguttata Barrera-Rodríguez and Ruíz-Carranza (Guayasamin et al. 2009). Hyalinobatrachium aureogut- 1989: 77. tatum is superficially similar to H. valeroi. Nevertheless, Hyalinobatrachium aureoguttatum — Ruíz-Carranza and Lynch 1991: in preservative H. aureoguttatum differs from H. valeroi 23. in having usually either well-defined rounded spots or Material examined. Table 1. elongate blotches of bright white on yellow background Restrepo et al. | Herpetofauna of the northern Andes of Colombia 209

Figure 2. Amphibian species present in Jaguas and San Carlos hydroelectric protected areas. A: Rheobates pseudopalmatus MHUAA 8491. B: Rhinella alata MHUAA 8486. C: Rhinella horribilis MHUAA 10722. D: Cochranella resplendens MHUAA 9540. E: Espadarana prosoblepon MHUAA 8493. F: Hyalinobatrachium aureogutattum MHUAA 7075. G: Hyalinobatrachium fleischmanni MHUAA 8176. H: Rulyrana susatamai MHUAA 10684. 210 Check List 13 (4) which correspond to clumps of iridophores (absent in H. in C. fitzingeri), and by having the posterior surfaces of valeroi). the thighs unicolor (with pale spots in C. fitzingeri). Hyalinobatrachium fleischmanni (Boettger, 1893): Pristimantis gaigei (Dunn, 1931): Figure 3E Figure 2 Lithodytes gaigei Dun 1931: 387. Hylella fleischmannBoettger 1893: 251. Pristimantis gaigeae — Hedges et al. 2008: 120. Hyalinobatrachium fleischmanni — Ruiz-Carranza and Lynch 1991: 24. Material examined. Table 1. Material examined. Table 1. Pristimantis gaigei is a small to medium-sized frog characterized by having a distinct tympanum; annulus Rulyrana susatamai (Ruíz-Carranza & Lynch, 1995): partially concealed dorsally and posteriorly by supra- Figure 2H Cochranella susatamai Ruíz-Carranza and Lynch 1995): 62. tympanic fold; skin of dorsal surfaces shagreened; skin Rulyrana susatamai — Guayasamin et al. 2009: 34. of venter smooth or feebly granular along lateral and Material examined. Table 1. posterior borders of ventral disk; discoidal folds and prominent arm slender (Lynch 1980). This species is eas- Sachatamia punctulata (Ruiz-Carranza & Lynch, 1995): ily distinguished from related species (i.e., P. penelopus) Figure 3A also distributed in the Magdalena river valley by having Cochranella punctulata Ruiz-Carranza and Lynch 1995: 4. smooth and dark gray ventral surface, a bright yellow Sachatamia punctulata — Guayasamin et al. 2009: 36. band on the forearm, and no calcar tubercle (in contrast Material examined. Table 1. with brown ventral surface with cream flecking, absence Sachatamia sp.: Figure 3B of yellow band and presence of calcar tubercle in P. Material examined. Table 1. penelopus). Craugastor metriosistus Ospina-Sarria, Angarita-Sierra Pristimantis jaguensis Rivera-Prieto, Rivera-Correa & & Pedroza-Banda, 2015: Figure 3C Daza, 2014: Figure 3F Craugastor metriosistus Ospina-Sarria et al. 2015: 165. Pristimantis jaguensis Rivera-Prieto, Rivera-Correa & Daza 2014: 225. Material examined. Table 1. Material examined. Table 1. A medium-sized frog characterized by having A moderated-sized frog characterized by a wide vari- extensive toe webbing and a uniformly reddish brown ation in color; absence of nuptial pads, discoidal fold and coloration on the posterior surfaces of the thighs. Crau- conical calcar tubercles; flanks and belly white to cream gastor metriosistus differ from C. longirostris and C. without blotches as well as iris yellow ocher to copper raniformis (the latter also distributed in the Magdalena with thick brown reticulation and cream sclera (Rivera river valley) in having the throat with median white gular et al. 2014). This species co-occurs in the Magdalena stripe and dark coloration on either side (throat with spot- river valley with Pristimantis penelopus and P. viejas ted pattern or short lines in C. longirostris and white to and may be distinguished from both by having smooth cream in C. raniformis), in having posterior surfaces of skin and lacking a calcar tubercle (dorsum with tubercles the thighs reddish brown (dark to reddish brown with including the heel in P. penelopus and P. viejas) and the small and conspicuous pale spots in C. raniformis), and concealed surfaces of the thighs yellow without blotches in having webbing extending to the proximal portion of (orange spots on the concealed surfaces of the limbs in the distal subarticular tubercle on the outer side of toe III P. viejas; yellow spots on the concealed surfaces of the (webbing extends to middle portion between basal and limbs, sometimes forming a reticle, in P. penelopus). distal subarticular tubercle on the outer side of toe III in Pristimantis penelopus (Lynch & Rueda-Almonacid, C. raniformis and C. longirostris; Ospina-Sarria et al. 1999): Figure 3G 2015). Eleutherodactylus penelopus Lynch and Rueda-Almonacid 1999: 310. Pristimantis fallax (Lynch & Rueda-Almonacid, 1999): Pristimantis penelopus — Hedges et al. 2008: 128. Figure 3D Material examined. Table 1. Eleutherodactylus fallax Lynch and Rueda-Almonacid 1999: 308. A moderated-sized frog with a shagreen dorsum; Pristimantis fallax — Hedges et al. 2008: 118. tubercles posteriorly and on flanks; no dorsolateral folds; Material examined. Table 1. skin of venter brown with cream flecking; discoidal Pristimantis fallax is a moderate size frog with a folds prominent; subanal tubercles only slightly larger shagreen dorsum; finger I longer than finger II; toeV than granules on underside of thighs; and iris copper slightly longer than toe III; absence of toe webbing and with black reticulation. Males have spots on concealed broad disks on outer fingers. The most distinctive feature surfaces of the thighs, a feature shared with Pristiman- of P. fallax is the coloration pattern of the throat consisting tis viejas; nevertheless, these spots are usually yellow in a lateral portion reticulated with gray to black, defin- (sometimes forming a reticle) in P. penelopus and orange ing cream central raphe, a feature shared with Craugastor in P. viejas. Pristimantis penelopus is also similar to P. fitzingeri. Although superficially resemblingC. fitzingeri, latidiscus from which it may distinguished in having a P. fallax is easily distinguished in having toe V slightly brown venter with cream flecking (in contrast to a cream longer than toe III (toe III slightly longer than toe V in C. venter with brown reticulation) (Lynch and Rueda-Almo- fitzingeri), absence of webbing on toes (basally webbed nacid 1999, Restrepo et al. 2017). Restrepo et al. | Herpetofauna of the northern Andes of Colombia 211

Figure 3. Amphibian species present in Jaguas and San Carlos hydroelectric protected areas. A: Sachatamia punctulata MHUAA 10689. B: Sachatamia sp. MHUAA 8205. C: Craugastor metriosistus MHUAA 10589. D: Pristimantis fallax MHUAA 10523. E: Pristimantis gaigei MHUAA 8497. F: Pristimantis jaguensis. G: Pristimantis penelopus MHUAA 8487. H: Pristimantis taeniatus MHUAA 8483. 212 Check List 13 (4)

Pristimantis taeniatus (Boulenger, 1912): Figure 3H Diasporus anthrax (Lynch, 2001): Figure 4E Eleutherodactylus taeniatus Boulenger 1912: 188. Eleutherodactylus anthrax Lynch 2001: 292. Pristimantis taeniatus — Hedges et al. 2008: 121. Diasporus anthrax — Hedges et al. 2008: 47. Material examined. Table 1. Material examined. Table 1. Small frog with skin of dorsum smooth; tympanum Pristimantis viejas (Lynch & Rueda-Almonacid, 1999): round and prominent; snout subacuminate in dorsal Figure 4A Eleutherodactylus viejas Lynch and Rueda-Almonacid 1999: 311. view, rounded in lateral view; all fingers and toes with Pristimantis viejas — Hedges et al. 2008: 121. expanded disks; pads triangular on toes III–IV; dorsum Material examined. Table 1. brown with darker blotches and venter dark brown with A small-sized frog with numerous non-conical whitish flecks. Diasporus anthrax differs from other tubercles on dorsum; absence of dorsolateral folds; finger species of the genus by its coloration which consist in I shorter than finger II; toe V very long; brown above a dark venter with white flecks and pale blotches (red in with darker brown markings; venter cream stippled with life) on the concealed surfaces of the limb (orange above, brown and posterior surfaces of the thighs brown with venter immaculate, and lacking distinctive coloration large cream spots. Superficially similar to Pristimantis marks on limbs in D. gularis; brown above, venter white taeniatus in lacking a canthal stripe, but readily distin- with brown marbling, and lacking distinctive coloration guished from that species in having orange spots on the marks on limbs in D. tinker) and the rounded disk pads concealed surfaces of the limbs (absent in P. taeniatus; and disk covers except for finger III and toes III–IV in Lynch and Rueda-Almonacid 1999). contrast with all fingers bearing narrow disks and disk cover of finger III bearing elongate papilla in D. tinker; Colostethus inguinalis (Cope, 1868): Figure 4B Prostherapis inguinalis Cope 1868: 10. unornamented disk covers and knobbed disk cover occur Colostethus inguinalis — Savage 1968: 745. in about equal frequency in D. gularis (Lynch 2001). Material examined. Table 1. Diasporus gularis (Boulenger, 1898): Figure 4F A moderate-sized dendrobatid with a pale oblique lat- Hylodes gularis Boulenger 1898: 121. eral stripe extending from groin only to around midbody; Diasporus gularis — Hedges et al. 2008: 47. pale ventrolateral stripes present; pale dorsolateral stripes Material examined. Table 1. absent and toes moderately webbed. Colostethus ingui- A small frog with short body; snout nearly truncate at nalis may be distinguished from C. aff. fraterdanieli (also tip when viewed from above, truncate in profile; nostrils distributed in the Magdalena river valley) by having an strongly projecting laterally; fingers very short, with lat- incomplete pale oblique lateral stripe (complete in C. aff. eral ridges, pointed disks, and very small basal webbings; fraterdanieli) and by the throat coloration of adults (black skin of upper surfaces smooth with a few short weak in adult males, white unpigmented or faintly pigmented glandular lines above shoulders and along sides. The best gray or brown in adult females in C. inguinalis; spotted diagnostic characters to recognize Diasporus gularis are in adult males and immaculate in adult females in C. aff. its very short hands and feet with pointed fingers and toes, fraterdanieli). It differs from C. pratti both in toe web- of which the entire terminal disk is dark gray, and the bing (absent in C. pratti) and adult male throat coloration enormous U-shaped gular pouch of the males (Cochran (pale gray or brown with irregular white spots, forming a and Goin 1970). Diasporus gularis is most readily dis- faint, mottled or reticulated pattern in C. pratti). tinguished from other Diasporus (i.e., D. anthrax and Colostethus aff. fraterdanieli: Figure 4C D. tinker) by its coloration which is orange above with Material examined. Table 1. a transverse dark bar between eyes and immaculate ven- A moderate-sized dendrobatid with dorsal skin texture ter (dorsum brown with darker blotches and venter dark granular posteriorly; pale dorsolateral stripe absent; pale brown with whitish flecks in D. anthrax; brown above oblique lateral stripe complete (extending from groin to with a tan interorbital bar, dorsolateral flank stripes and eye); pale ventrolateral stripe present; coloration pattern venter white with brown marbling in D. tinker; Lynch of throat spotted in males, immaculate in females (Tolosa 2001). et al. 2014). Colostethus aff. fraterdanieli differs from C. Hemiphractus fasciatus Peters, 1862: Figure 4G fraterdanieli in having a pale ventrolateral stripe (absent Hemiphractus fasciatus Peters 1862: 169. in C. fraterdanieli) and a continuous dark band running Boana boans (Linnaeus, 1758): Figure 4H along the lateral surfaces of body and head (discontinu- Rana boans Linnaeus 1758: 213. ous in C. fraterdanieli). It may be distinguished from C. Hypsiboas boans — Faivovich et al. 2005: 89. inguinalis by having a complete oblique lateral stripe Boana boans — Dubois 2017: 28. (partial in C. inguinalis) and from C. pratti by lacking Material examined. Table 1. pale dorsolateral stripe (present in C. pratti). Boana xerophylla (Duméril & Bibron, 1841): Figure 5A Dendrobates truncatus (Cope, 1861): Figure 4D Hyla xerophylla Duméril and Bibron 1841: 549. Phyllobates truncatus Cope 1861a: 372. Hypsiboas xerophyllus — Orrico et al. 2017: 107. Dendrobates truncatus — Cope 1867: 197. Boana xerophylla — Dubois 2017: 28. Material examined. Table 1. Material examined. Table 1. Restrepo et al. | Herpetofauna of the northern Andes of Colombia 213

Figure 4. Amphibian species present in Jaguas and San Carlos hydroelectric protected areas. A: Pristimantis viejas MHUAA 9742. B: Colos- tethus inguinalis MHUAA. 10645. C: Colostethus aff. fraterdanieli MHUAA 8140. D: Dendrobates truncatus MHUAA 9740. E: Diasporus anthrax MHUAA 7237. F: Diasporus gularis MHUAA 8496. G: Hemiphractus fasciatus. H: Boana boans MHUAA 10564. 214 Check List 13 (4)

This species was recently separated from Boana crepi- Hyloscirtus palmeri (Boulenger, 1908): Figure 5F tans (Orrico et al. 2017). A large-sized tree frog having its Hyla palmeri Boulenger 1908: 515. dorsal surfaces pinkish tan with some darker brown spots, Hyloscirtus palmeri — Faivovich et al. 2005: 85. blotches or other markings; dark bars on the upper sur- Material examined. Table 1. faces of the thighs extending onto the rear surfaces of the Phyllomedusa venusta Duellman & Trueb, 1967: Figure thighs; ventral surface orange, except for the throat and 5G chest, which are white with some brown flecking. Boana Phyllomedusa venusta Duellman and Trueb 1967: 128. xerophylla may be confused with B. boans and B. pugnax Material examined. Table 1. (species distributed in the Magdalena river valley) due to Scinax sp.: Figure 5H similarities in body size and coloration. However, it may Material examined. Table 1. be distinguished by its horizontal pupil and its iris that is Our Scinax sp. corresponds to Scinax “A”, an unde- pale gray at the center and greenish yellow at the edges scribed species proposed by Nieto-Castro (1999) using (in contrast to a rhomboid pupil with an iris that is black at morphological traits, morphometric and chromatic analy- the center and light orange at the edges in B. boans, and a ses. From that study, the name began to be used to refer to horizontal pupil with an iris that is black at the center and populations of Scinax distributed in Antioquia and Cun- greenish in the upper portion in B. pugnax). dinamarca, characterized by having a golden color with a Dendropsophus ebraccatus (Cope, 1874): Figure 5B dorsal spotted pattern. Hyla ebraccata Cope 1874: 69. Dendropsophus ebraccatus — Faivovich et al. 2005: 91. Scinax rostratus (Peters, 1863): Figure 6A Hyla rostrata Peters 1863a: 466. Material examined. Table 1. Scinax rostratus — Köhler and Böhme 1996: 139. Dendropsophus microcephalus (Cope, 1886): Figure 5C Material examined. Table 1. Hyla microcephala Cope 1886: 281. A moderate-sized hylid frog characterized by a con- Dendropsophus microcephalus — Faivovich et al. 2005: 92. spicuous acuminate and projecting snout in dorsal view, Dendropsophus norandinus Rivera-Correa & Guitérrez- protruding in lateral view (Duellman 1970). Scinax Cárdenas, 2012: Figure 5D rostratus is readily distinguished from Scinax ruber and Dendropsophus norandinus Rivera-Correa & Guitérrez-Cárdenas 2012: Smilisca sila (both species occurring in the Magdalena 51. river valley) by the combination of its characteristic snout Material examined. Table 1. and coloration pattern (surfaces of thighs yellow or pale A small hylid frog with a snout that is short, rounded orange with black or dark brown markings in S. rostratus; in dorsal view and slightly rounded in lateral view; head distinct black and white markings on posterior thighs in wider than long; skin on dorsal surfaces smooth, with S. ruber; blue spots on flanks and posterior surfaces of the some minute scattered tubercles; axillary membrane thighs in S. sila). present and developed; dorsum reddish brown in females Scinax ruber (Laurenti, 1768): Figure 6B and cream or silver gray in males (Rivera-Correa and Hyla rubra Laurenti 1768: 35. Guitérrez-Cárdenas 2012). Dendropsophus norandinus Scinax ruber — Kölher and Böhme 1996: 139. may be distinguished from D. bogerti by having the head Material examined. Table 1. wider than long (longer than wide in D. bogerti) and by A medium-sized frog with toes webbed except for having finger webbing that extends to the base ofthe the innermost; dorsal surfaces generally olive brown disks of toes I, II, and III (finger webbing absent in D. or orange-brown; distinct black and white markings on bogerti), from D. subocularis by lacking light bands con- posterior surfaces of thighs. Related species which may necting eye to edge of lips, and from D. microcephalus be confused with Scinax ruber are S. boulengeri and S. by its webbed fingers (fingers free in D. microcephalus). rostratus (also distributed in the Magdalena river val- ley) from both of these it differs by its smaller size and Dendropsophus subocularis (Dunn, 1934): Figure 5E the presence of black markings on the thighs in form of Hyla subocularis Dunn 1934: 2. Dendropsophus subocularis — Faivovich et al. 2005: 93. vermiculations instead of broad and vertical dark bands Material examined. Table 1. (Cochran and Goin 1970). Scinax ruber is also similar to A small hylid frog with short snout; canthus rostralis Smilisca phaeota from which it differs by lacking green very marked; fingers webbed at base about toes I–III; coloration in the rostral region, and to Smilisca sila from toes webbed nearly to disks of III and V; skin of dorsum which it differs by the absence of blue spots on the flanks smooth and skin of vent rugose (Dunn 1934). Dendrop- and posterior surfaces of the thighs. sophus subocularis co-occurs with D. microcephalus and Smilisca phaeota (Cope, 1862): Figure 6C D. norandinus in the Magdalena river valley. It may be Hyla phaeota Cope 1862: 358. distinguished from both by the presence of 2 light bands Smilisca phaeota — Starrett 1960: 303. from the eye to the edge of the lips, enclosing a bar of Material examined. Table 1. dark between them (absent in the latter 2 species), and Engystomops pustulosus (Cope, 1864): Figure 6D from D. microcephalus by its webbed fingers (fingers free Paludicola pustulosa Cope 1864: 180. in D. microcephalus). Engystomops pustulosus — Boulenger 1882: 276. Restrepo et al. | Herpetofauna of the northern Andes of Colombia 215

Figure 5. Amphibian species present in Jaguas and San Carlos hydroelectric protected areas. A: Boana xerophyllus MHUAA 10567. B: Dendropsophus ebraccatus MHUAA 10692. C: Dendropsophus microcephalus MHUAA 8667. D: Dendropsophus norandinus MHUAA 8589. E: Dendropsophus subocularis MHUAA 8490. F: Hyloscirtus palmeri MHUAA 9504. G: Phyllomedusa venusta MHUAA 10581. H: Scinax sp. MHUAA 8228. 216 Check List 13 (4)

Figure 6. Amphibian species present in Jaguas and San Carlos hydroelectric protected areas. A: Scinax rostratus MHUAA 7233. B: Scinax ruber MHUAA 9500. C: Smilisca phaeota MHUAA 9503. D: Engystomops pustulosus MHUAA 10602. E: Leptodactylus colombiensis MHUAA 8818. F: Leptodactylus fuscus MHUAA 10598. G: Leptodactylus savagei MHUAA 9768. H: Bolitoglossa ramosi. Restrepo et al. | Herpetofauna of the northern Andes of Colombia 217

Figure 7. Amphibian species present in Jaguas and San Carlos hydroelectric protected areas. A: Caecilia thompsoni MHUAA 7192. B: Microcaecilia pricei MHUA-A 10728. C: Epicrionops parkeri MHUA- A 10720.

Physalaemus pustulosus — Lynch 1970: 488. Microcaecilia pricei — Wilkinson, O’Connor and Nassbaum 2013: 7. Material examined. Table 1. Material examined. Table 1. Leptodactylus colombiensis Heyer, 1994: Figure 6E Epicrionops parkeri (Dunn, 1942): Figure 7C Leptodactylus colombiensis Heyer 1994: 82. Rhinatrema parkeri Dunn 1942: 458. Material examined. Table 1. Epicrionops parkeri — Taylor 1968: 216. Material examined. Table 1. Leptodactylus fuscus (Schneider, 1799): Figure 6F A bicolored caecilian with well-developed tail; lat- Rana fusca Schneider 1799: 130. Leptodactylus fuscus — Heyer 1968: 160. eral stripe yellow, cream or white from head or second Material examined. Table 1. collar onto tail; stripe ending at posterior level of vent; subcaudal region dark lilac to blackish (Taylor 1968). Leptodactylus savagei Heyer, 2005: Figure 6G Epicrionops parkeri differs from E. bicolor (a species Leptodactylus savagei Heyer 2005: 330. with similar coloration pattern distributed in Colombia) Bolitoglossa ramosi Brame & Wake, 1972: Figure 6H by having a larger body length and 198 primary annuli (in Bolitoglossa ramosi Brame and Wake 1972: 9. contrast to 117–135 primary annuli in E. bicolor). Material examined. Table 1. Caecilia thompsoni Boulenger, 1902: Figure 7A Reptiles Caecilia thompsoni Boulenger 1902: 152. We report 58 reptile species comprising 47 genera in 16 Material examined. Table 1. families (Table 2). The protected area of Jaguas hydro- A large caecilian with generally cylindrical body; eye electric project has 38 species from 14 families, where in an open orbit and tentacular aperture below nostril, the most speciose families were Dipsadidae and Colub- close to the anterior portion of the face (Lynch 1999). ridae with 10 and 8 species, respectively (26 and 21%). Caecilia thompsoni is readily distinguished from C. sub- One species (Anolis sulcifrons Daudin, 1802) is endemic nigricans (another species known from Magdalena river to Colombia and only 10 (26%) have been evaluated by valley) by having a longer body (up to 1.5 m in C. thomp- the IUCN, 1 listed as Near Threatened and 9 as Least soni; 0.35–0.37 m in C. subnigricans) and more primary Concern (Table 4). In contrast, in San Carlos we found (187–240 in C. thompsoni; 145–169 in C. subnigricans) 57 reptile species comprising 16 families. Again, the and secondary annuli (26–30 in C. thompsoni; 6–27 in C. most speciose families at this site were Dipsadidae with subnigricans) 15 species (26%) and Colubridae with 11 species (19%). Microcaecilia pricei (Dunn, 1944): Figure 7B Two species, Anolis sulcifrons Daudin, 1802 and Ninia Gymnopis pricei Dunn 1944: 473. teresitae Angarita-Sierra & Lynch, 2017 are endemic to 218 Check List 13 (4)

Colombia (4%) and 16 have been evaluated by the IUCN the fore and hind limbs (Oftedal 1974, Savage 2002). (2016), all of them categorized as Least Concern. Jaguas Usually, anterior most spots are bordered by black, and San Carlos localities shared 36 reptile species. forming ocelli. Although there is no evidence of Anadia ocellata for the Magdalena river valley, we assigned the Basiliscus galeritus Duméril, 1851: Figure 8A Basiliscus galeritus Duméril 1851: 61. specimen MHUAR 12694 to Anadia gr. ocellata because Material examined. Table 2. its dorsal coloration and 55 transversal rows in contrast Adults of this large semiaquatic lizard are easily to the 44–48 dorsal rows found in A. rhombifera, another recognized by their gray to green color and the presence species present in the Magdalena river valley (Oftedal of a single, rounded head crest that is larger in males. 1974, Savage 2002). Similar species that are found sympatric with B. galeri- Cercosaura sp.: Figure 8H tus are Corytophanes cristatus and Basiliscus basiliscus. Material examined. Table 2. The first is usually brownish and has 2 head crests that converge posteriorly. Basiliscus basiliscus has a brown Echinosaura palmeri Boulenger, 1911: Figure 9A Echinosaura palmeri Boulenger 1911: 23. or gray coloration and a single pointed head crest that is Material examined. Table 2. larger in males. Males of B. basiliscus also have a large crest in the back that resembles a fin (Maturana 1962; Loxopholis rugiceps Cope, 1869 Savage 2002; Köhler 2008). Loxopholis rugiceps Cope 1869: 305. Material examined. Table 2. Corytophanes cristatus (Merrem, 1820): Figure 8B Agama cristata Merrem 1820: 50. Loxopholis southi (Ruthven & Gaige, 1924): Figure 9B Corytophanes cristatus —Boie in Schlegel 1827: 290. Leposoma southi Ruthven and Gaige 1924: 1. Material examined. Table 2. Loxopholis southi — Goicoechea et al. 2016: 32. Material examined. Table 2. Anolis auratus Daudin, 1802: Figure 8C This species is easily differentiated from other gym- Anolis auratus Daudin 1802: 89. nophthalmid lizards in the region by its strongly keeled Material examined. Table 2. head and dorsal scales and by having the ventral scales Anolis fuscoauratus D’Orbigny, 1837 large, keeled, mucronate, and quadrangular to rectangu- Anolis fuscoauratus D’Orbigny in Duméril and Bibron 1837: 110. lar. It differs from Loxopholis rugiceps, the only other Material examined. Table 2. Loxopholis in trans-Andean Colombia, by having the frontonasal divided, instead of single (Köhler 2008). Anolis aff. granuliceps: Figure 8D Material examined. Table 2. Ptychoglossus sp.: Figure 9C Material examined. Table 2. Anolis sulcifrons Cope, 1899: Figure 8E Anolis sulcifrons Cope 1899: 6. Thecadactylus rapicauda (Houttuyn, 1782): Figure 9D Material examined. Table 2. Gekko rapicauda Houttuyn 1782: 323. Thecadactylus rapicauda — Boulenger 1885: 111. Anolis tropidogaster Hallowell, 1956: Figure 8F Material examined. Table 2. Anolis tropidogaster Hallowell 1956: 224. Polychrus gutturosus Berthold, 1846: Figure 9E Anolis vittigerus Cope, 1862 Polychrus gutturosus Berthold 1846: 11. Anolis vittigerus Cope 1862: 179. This large blunt-headed lizard is immediately recog- Material examined. Table 2. nizable by its extremely long round tail that is at least Anadia gr. ocellata Gray, 1845: Figure 8G twice the snouth–vent length, its robust body and rela- Material examined. Table 2. tively thin legs, and the presence of a large subtympanic This small lizard may be distinguished from most scale (Peters and Donoso-Barros 1970, Savage 2002). other mid- to low-elevation gymnophthalmid lizards in In addition, it lacks toe pads, nuchal, dorsal and caudal having an acuminate snouth and elongate body in com- crests (Peters and Donoso-Barros 1970, Savage 2002). bination with smooth head and dorsal scales. Its color It differs from Polychrus marmoratus, the other species pattern includes a series of lateral white spots between that could be sympatric, because the latter has a row of

Table 2. Voucher list of reptiles of Jaguas and San Carlos hydroelectric compensation areas with geographic coordinates.

Taxon Voucher Latitude Longitude Taxon Voucher Latitude Longitude REPTILIA MHUA-R 13006 6.29922 -74.96633 SQUAMATA MHUA-R 13173 6.3515 -74.99528 SAURIA Corytophanes cristatus MHUA-R 12468 6.21854 -74.83727 Corytophanidae MHUA-R 12429 6.24537 -74.86961 Basiliscus galeritus MHUA-R 13010 6.277 -74.93104 MHUA-R 13015 6.28288 -74.92422 MHUA-R 13009 6.2832 -74.92413 MHUA-R 13191 6.19012 -74.80477 MHUA-R 13016 6.29256 -75.04247 Continued Restrepo et al. | Herpetofauna of the northern Andes of Colombia 219

Table 2. Continued.

Taxon Voucher Latitude Longitude Taxon Voucher Latitude Longitude Dactyloidae MHUA-R 13174 6.36704 -75.02715 Anolis auratus MHUA-R 12432 6.2199 -74.84463 Amphisbaenidae MHUA-R 13216 6.20643 -74.83891 Amphisbaena varia MHUA-R 12276 6.19073 -74.80437 MHUA-R 13217 6.20643 -74.83891 Colubridae MHUA-R 13218 6.20643 -74.83891 Chironius grandisquamis MHUA-R 15011 6.36728 -75.02697 Anolis fuscoauratus MHUA-R 12433 6.21537 -74.83533 Dendrophidion MHUA-R 14764 6.19073 -74.80437 Anolis aff. granuliceps MHUA-R 12726 6.19062 -74.80474 percarinatum MHUA-R 12792 6.19086 -74.80454 Drymobius rhombifer MHUA-R 15012 6.39058 -75.03704 MHUA-R 12514 6.1957 -74.8167 Mastigodryas boddaerti MHUA-R 15054 6.19062 -74.80474 MHUA-R 12482 6.21768 -74.83748 MHUA-R 15078 6.28296 -74.92384 MHUA-R 13013 6.32754 -75.01631 MHUA-R 14866 6.22879 -74.87466 MHUA-R 13011 6.32781 -75.01151 Phrynonax poecilonotus MHUA-R 15013 6.19675 -74.80444 MHUA-R 13014 6.32791 -75.01171 Scaphiodontophis MHUA-R 14871 6.2189 -74.8477 MHUA-R 12725 6.36725 -75.02723 venustissimus MHUA-R 12513 6.3693 -75.0252 Tantilla melanocephala MHUA-R 14855 6.21854 -74.83727 MHUA-R 12445 6.37236 -75.04374 MHUA-R 14842 6.22148 -74.8418 MHUA-R 12745 6.38528 -75.02158 Dipsadidae MHUA-R 12695 6.38871 -75.02335 Atractus occipitoalbus MHUA-R 15131 6.36767 -75.02635 MHUA-R 12746 6.39128 -75.0364 Coniophanes fissidens MHUA-R 14865 6.21691 -74.81272 MHUA-R 12438 6.39677 -75.01162 MHUA-R 14879 6.2189 -74.8477 MHUA-R 12436 6.40714 -75.02618 MHUA-R 14846 6.3852 -74.99314 MHUA-R 12442 6.21611 -75.00078 MHUA-R 15133 6.20908 -74.85436 MHUA-R 12441 6.37088 -75.04567 Erythrolamprus epinephelus MHUA-R 14841 6.20874 -74.81525 MHUA-R 12440 6.37187 -75.04636 Erythrolamprus MHUA-R 14826 6.349358 -75.002699 MHUA-R 12435 6.37588 -75.03902 pseudocorallus MHUA-R 12019 6.8906 -75.5067 MHUA-R 14856 6.20716 -74.81999 MHUA-R 12393 7.393528 -75.036633 Imantodes cenchoa MHUA-R 14978 6.36725 -75.02723 MHUA-R 13211 6.39912 -75.02728 MHUA-R 15134 6.36704 -75.02715 MHUA-R 13226 6.1401 -75.06688 Leptodeira septentrionalis MHUA-R 15077 6.32938 -75.01601 MHUA-R 13227 6.14229 -75.06267 MHUA-R 14597 6.348145 -75.005172 Anolis sulcifrons MHUA-R 12696 6.19055 -74.8048 Ninia atrata MHUA-R 14847 6.2199 -74.84463 MHUA-R 12431 6.20801 -74.8157 MHUA-A 15130 6.23974 -74.8688 MHUA-R 12434 6.21146 -74.82359 Ninia teresitae MHUA-R 14860 6.21626 -74.83435 MHUA-R 12464 6.21872 -74.84776 Oxyrhopus petolarius MHUA-R 14843 6.21146 -74.82359 MHUA-R 12443 6.24739 -74.86933 MHUA-R 14693 6.8906 -75.5067 Anolis vittigerus MHUA-R 12512 6.1957 -74.8167 MHUA-R 15150 6.38927 -74.99146 Gymnophthalmidae MHUA-R 15151 6.25277 -74.82888 Anadia gr. ocellata MHUA-R 12694 6.1957 -74.8167 Rhadinaea decorata MHUA-R 14859 6.2167 -74.86468 Cercosaura sp. MHUA-R 12762 6.409987 -75.14157 Sibon nebulatus MHUA-R 14857 6.20716 -74.81999 MHUA-R 13221 6.287778 -74.828611 MHUA-R 15139 6.38747 -74.99274 Echinosaura palmeri MHUA-R 13212 6.3995 -75.02823 MHUA-R 15149 6.38927 -74.99151 MHUA-R 13213 6.3994 -74.0284 MHUA-R 15154 6.15764 -75.0416 MHUA-R 13223 6.15816 -75.04485 Siphlophis compressus MHUA-R 14733 6.1738 -74.7513 MHUA-R 13224 6.15813 -75.04476 Urotheca fulviceps MHUA-R 14979 6.19062 -74.80474 MHUA-R 13225 6.1584 -75.04494 MHUA-R 15055 6.19062 -74.80474 Loxopholis rucigeps MHUA-R 12107 6.2247 -74.8477 MHUA-R 15132 6.37516 -74.98231 Loxopholis southi MHUA-R 12430 6.20671 -74.855 Elapidae MHUA-R 12465 6.21841 -74.84771 Micrurus dumerilii MHUA-R 15137 6.19032 -74.80515 MHUA-R 12105 6.2275 -74.8525 MHUA-R 15138 6.19032 -74.80515 MHUA-R 12466 6.22787 -74.872264 Micrurus mipartitus MHUA-R 14958 6.25141 -74.82854 MHUA-R 12106 6.2275 -74.8525 MHUA-R 15152 6.19228 -74.80225 Ptychoglossus sp. MHUA-R 12602 6.36665 -75.02802 Leptotyphlopidae Phyllodactylidae Trilepida macrolepis MHUA-R 14858 6.21734 -74.81262 Thecadactylus rapicauda MHUA-R 12601 6.19132 -74.80448 Viperidae Sphaerodactylidae Bothrops asper MHUA-R 14039 6.20453 -74.8949 Gonatodes albogularis MHUA-R 12423 6.231083 -74.876611 MHUA-R 14691 6.8906 -75.5067 Lepidoblepharis MHUA-R 12467 6.2072 -74.81239 Porthidium lansbergii MHUA-R 14054 6.20453 -74.8949 xanthostigma MHUA-R 12471 6.2167 -74.86468 MHUA-R 14516 6.224778 -74.847778 MHUA-R 12422 6.231083 -74.876611 MHUA-R 15079 6.27689 -74.93108 MHUA-R 13012 6.29719 -74.91953 MHUA-R 14514 6.352361 -75.012139 MHUA-R 13008 6.29826 -74.91976 MHUA-R 14515 6.354778 -75.008278 MHUA-R 12698 6.39202 -75.0365 MHUA-R 14956 6.38553 -75.0218 Sphaerodactylus lineolatus MHUA-R 12470 6.18822 -74.8045 MHUA-R 14694 6.511907 -74.910799 MHUA-R 12728 6.19062 -74.80474 MHUA-R 14513 6.399989 -75.010364 MHUA-R 12469 6.21856 -74.83739 TESTUDINES Teiidae Kinosternidae Holcosus festivus MHUA-R 12444 6.21537 -74.83533 Kinosternon leucostomum MHUA-R 18526 6.38871 -75.02555 MHUA-R 12018 6.8906 -75.5067 MHUA-R 18310 6.1897 -74.9969 220 Check List 13 (4)

Figure 8. Reptile species present in Jaguas and San Carlos hydroelectric protected areas. A: Basiliscus galeritus MHUA-R 13009. B: Coryto- phanes cristatus MHUA-R 13015. C: Anolis auratus MHUA-R 13217. D: Anolis aff.granuliceps MHUA-R 13011. E: Anolis sulcifrons MHUA-R 12464. F: Anolis tropidogaster. G: Anadia gr. ocellata MHUA-R 12694. H: Cercosaura sp. MHUA-R 13162. Restrepo et al. | Herpetofauna of the northern Andes of Colombia 221

Figure 9. Reptile species present in Jaguas and San Carlos hydroelectric protected areas. A: Echinosaura palmeri MHUA-R 13158. B: Loxopho- lis southi MHUA-R 12466. C: Ptychoglossus sp. MHUA-R 12602. D: Thecadactylus rapicauda MHUA-R 12601. E: Polychrus gutturosus. F: Marisora sp. MHUA-R 13210. G: Gonatodes albogularis MHUA-R 12424. H: Lepidoblepharis xanthostigma MHUA-R 13012. 222 Check List 13 (4)

Figure 10. Reptile species present in Jaguas and San Carlos hydroelectric protected areas. A: Sphaerodactylus lineolatus MHUA-R 12470. B: Ameiva sp. C: Cnemidophorus lemniscatus. D: Holcosus festivus MHUA-R 13174. E: Amphisbaena varia. F: Boa constrictor. G: Chironius exole- tus. H: Chironius grandisquamis MHUA-R 15011. Restrepo et al. | Herpetofauna of the northern Andes of Colombia 223 enlarged scales, forming a crest, in the mental and gular separated from its sympatric congeners by having 10 rows regions, which is absent in P. gutturosus (Peters and of dorsal scales with the paravertebral scales keeled (12 Donoso-Barros 1970). rows in Chironius exoletus and C. carinatus), by having its dorsum black, or brown with transversal bands (dor- Gonatodes albogularis (Duméril & Bibron, 1836): sally green in C. exoletus and yellowish or olive ground Figure 9G Gymnodactylus albogularis Duméril and Bibron 1836: 415. color with 2 broad reddish brown stripes in C. carinatus). Gonatodes albogularis — Boulenger 1885: 59. Dendrophidion percarinatum (Cope, 1893): Figure 11A Material examined. Table 2. Drymobius percarinatum Cope 1893: 344. Lepidoblepharis xanthostigma (Noble, 1916): Figure 9H Dendrophidion percarinatum — Smith 1941: 73 Lathrogecko xanthostigma Noble 1916: 87. Material examined. Table 2 Lepidoblepharis xanthostigma — Vanzolini 1953: 264. Drymobius rhombifer (Günther, 1860): Figure 11B Material examined. Table 2. Coryphodon rhombifer Günther 1860: 236. This speices differs from other Lepidoblepharis Drymobius rhombifer — Boulenger 1894: 14. species by having granular dorsals (cycloid in L. sanc- Material examined. Table 2. taemartae), 12–14 lamellae under the fourth toe (<10 in L. victormartinezi and L. emberawoundule; Batista et al. Leptophis ahaetulla (Linnaeus, 1758): Figure 11C Coluber ahaetulla Linnaeus 1758: 225. 2015), median subcaudal scales more than twice as wide Leptophis ahaetulla — Bell 1825: 328. as the laterally adjacent subcaudals (less than 2 times in L. rufigularis; Batista et al. 2015). Mastigodryas boddaerti (Sentzen, 1796): Figure 11D Coluber boddaerti Sentzen 1796: 59. Sphaerodactylus lineolatus Lichtenstein & von Martens Mastigodryas boddaerti — Montingelli et al. 2011: 189. 1856: Figure 10A Material examined. Table 2. Sphaerodactylus lineolatus Lichtenstein & von Martens in Lichtenstein 1856: 6. Oxybelis aeneus (Wagler, 1824): Figure 11E Material examined. Table 2. Dryinus aeneus Wagler in Spix 1824: 12. Oxybelis aeneus — Duméril and Bibron 1854: 819. Cnemidophorus lemniscatus (Linnaeus, 1758): Figure 10C Phrynonax poecilonotus (Günther, 1858): Figure 11F Spilotes poecilonotus Günther 1858: 100. Lacerta lemniscata Linnaeus 1758: 209. Phrynonax poecilonotus — Boulenger 1894: 20. Cnemidophorus lemniscatus — Duméril and Bibron 1839: 129. Material examined. Table 2. Holcosus festivus (Lichtenstein, 1856): Figure 10D Adults have either red or yellow ground color, with Cnemidophorus festivus Lichtenstein 1856: 13. a variable dorsal pattern that may consist of black trans- Holcosus festivus Harvey et al. 2012: 118. versal bands or several black dots, mainly located in the Material examined. Table 2. posterior region of the scales. Other than coloration, Amphisbaena varia Laurenti, 1768: Figure 10E the species differs from other sympatric snakes by hav- Amphisbaena varia Laurenti 1768: 66. ing 21–25 dorsal rows at midbody, with the uppermost Material examined. Table 2. (3–19) rows keeled (Savage 2002). Some authors rec- This legless lizard is distinguished by its bullet-like ognize an additional species in trans-Andean Colombia, head and rectangular body scales arranged in trans- Phrynonax shropshirei (Taylor 1954, Peters et al. 1970, versal rings (annuli). It has a whitish or pink ground Pérez-Santos and Moreno 1988), but according to Sav- with a large amount of irregular brown blotches, often age (2002), it seems to be a confusion given the wide obscuring ground color (Vanzolini 2002). Amphisbaena color variation in P. poecilonotus. Likewise, the profound varia has traditionally been considered a subspecies of genetic distinction between P. poecilonotus and P. polyl- A. fuliginosa (Vanzolini 1951, 2002, Köhler 2008), but epis (Peters, 1876) revealed by Jadin et al. (2014), who Gans (2005) elevated it to species level without further accordingly resurrected the latter species, does apparently explanation. not correspond to any morphological distinction. Since Boa constrictor Linnaeus, 1758: Figure 10F neither Jadin et al. (2014) nor the original description Boa constrictor Linnaeus 1758: 215. (Peters 1876) offer any morphological characters to dis- criminate between these 2 species, we must assume such Chironius exoletus (Linnaeus, 1758): Figure 10G Coluber exoletus Linnaeus 1758: 223. distinctions to be obscured by the exuberant variability Chironius exoletus — Hoge, Romano and Cordeiro 1978: 41. in Phrynonax for now. Unable to answer the interesting question to which species our specimen actually pertains, Chironius grandisquamis (Peters, 1869): Figure 10H at this point we choose a conservative approach and ten- Spilotes grandisquamis Peters 1869: 451. Chironius grandisquamis— Savage 2002: 650. tatively assign it to P. poecilonotus. Material examined. Table 2. Scaphiodontophis venustissimus (Günther, 1893): This large species has a black or dark brown ground Figure 11G color and may have reddish or brown transversal bands Henicognathus venustissimus Günther 1893: 144. (Dixon et al. 1993, Savage 2002). This species may be Scaphiodontophis venustissimus — Taylor and Smith 1943: 309. 224 Check List 13 (4)

Figure 11. Amphibian species present in Jaguas and San Carlos hydroelectric protected areas. A: Dendrophidion percarinatum. B: Drymobius rhombifer MHUA-R 15012. C: Leptophis ahaetulla. D: Mastigodryas boddaerti MHUA-R 15054. E: Oxybelis aeneus. F: Phrynonax poecilonotus MHUA-R 15013. G: Scaphiodontophis venustissimus MHUA-R 14871. H: Spilotes pullatus. Restrepo et al. | Herpetofauna of the northern Andes of Colombia 225

Material examined. Table 2. Leptodeira septentrionalis Kennicott in Baird 1859: 16. Material examined. Table 2. Spilotes pullatus (Linnaeus, 1758): Figure 11H Coluber pullatus Linnaeus 1758: 225. Ninia atrata (Hallowell, 1845): Figure 13B Spilotes pullatus — Wagler 1830: 23. Coluber atratus Hallowell 1845: 245. This large snake reaches up to 2.5 m long. It differs Ninia atrata — Cope 1860: 340. from other snake species in the area by having a dorsal Material examined. Table 2. pattern of transversal yellow and black bands. Often, the Ninia tereseitae Angarita-Sierra & Lynch, 2017: Figure posterior portion of the body is completely black (Savage 13C 2002). It also differs from similar species in the region by Ninia tereseitae Angarita-Sierra and Lynch (2017): 478. having 14–18 dorsal rows of scales at the midbody with Material examined. Table 2. the uppermost keeled (Savage 2002). Oxyrhopus petolarius (Linnaeus, 1758): Figure 13D Tantilla melanocephala (Linnaeus, 1758): Figure 12A Coluber petola Linnaeus 1758: 225. Coluber melanocephalus Linnaeus 1758: 218. Oxyrhopus petolarius — Duméril and Bibron 1854: 1033. Tantilla melanocephala — Wilson and Mena 1980: 6. Material examined. Table 2. Material examined. Table Appendix 2. Rhadinaea decorata (Günther, 1858): Figure 13E Atractus occipitoalbus (Jan, 1862): Figure 12B Coronella decorata Günther 1858: 35. Rhabdosoma occipitualbum Jan 1862: 16. Rhadinaea decorata — Taylor 1949: 196. Atractus occipitualbus — Boulenger 1894: 310. Material examined. Table 2. Material examined. Table 2. This small, brown snake has a white, black-bordered Clelia clelia (Daudin, 1803): Figure 12C line that goes from the postoculars through the fifth and Coluber clelia Daudin 1803: 330. sixth dorsal rows to the tip of the tail. The vertebral Clelia clelia — Zaher 1996: 301. region is a little paler than below the lines. Specimens of Coniophanes fissidens (Günther, 1858): Figure 12D Rhadinaea decorata often have a subpreocular scale and Coronella fissidens Günther 1858: 36. 110–134 ventral scales (Myers 1974). The most similar Coniophanes fissidens —Pérez-Santos and Moreno 1988: 127. species in the region is Urotheca fulviceps, which has a Material examined. Table 2. reddish or purple head band, a pale line running along Erythrolamprus epinephelus (Cope, 1862): Figure 12E the first and second dorsal scale rows and more than 136 Liophis epinephalus Cope 1862: 78. ventrals. Erythrolamprus epinephelus — Grazziotin et al. 2012: 21. Sibon nebulatus (Linnaeus, 1758): Figure 13F Material examined. Table 2. Coluber sibon Linnaeus 1758: 222. Erythrolamprus melanotus (Shaw, 1802): Figure 12F Sibon nebulatus — Fitzinger 1826: 60. Coluber melanotus Shaw 1802: 534. Material examined. Table 2. Erythrolamprus melanotus — Grazziotin et al. 2012: 21. This species has a pattern of gray, white and black This medium-sized species may be recognized by bands. Gray bands are larger and with dispersed small having 17-17-15 smooth dorsal scale rows. Its dorsal black blotches, followed by a thin white band and then ground color is light brown and yellowish, with 3 dorsal by a medium-sized black band (Savage 2002). Ventrally, dark brown lines that extend from the neck to the tail. The ground is white with black blotches or bands. Sibon 2 more lateral lines, running along the fourth and fifth nebulatus has a large head and eyes like species of the dorsal rows, begin as series of points and the vertebral genus Imantodes, but it can be easily recognized by its line one is about 5 scale rows wide (Dixon and Michaud color pattern (Savage 2002). 1992). Ventral coloration is white or light yellow (Dixon Siphlophis compressus (Daudin, 1803): Figure 13G and Michaud 1992). The most similar sympatric species Coluber compressus Daudin 1803: 247. is Erythrolamprus epinephelus, which differs in having a Siphlophis compressus — Zaher and Prudente 1999: 699. grayish ground color with mainly transversal blotches or Material examined. Table 2. bands, 2 lateral dark lines that begin only in the posterior portion of the body extending to the tail, and frequently Urotheca fulviceps (Cope, 1886): Figure 13H a reddish to yellowish venter with black blotches (Dixon Rhadinaea fulviceps Cope 1886: 279. Urotheca fulviceps — Savage and Crother 1989: 343. 1983). Material examined. Table 2. Erythrolamprus pseudocorallus Roze, 1959: Figure 12G Erythrolamprus pseudocorallus Roze 1959: 530. Xenodon rabdocephalus (Wied-Neuwied, 1824): Material examined. Table 2. Figure 14A Coluber rabdocephalus Wied-Neuwied 1824: 668. Imantodes cenchoa (Linnaeus, 1758): Figure 12H Xenodon rabdocephalus — Duméril et al. 1854: 758. Coluber cenchoa Linnaeus 1758: 226. Micrurus dumerilii Jan, 1858: Figure 14B Material examined. Table 2. Micrurus dumerilii Jan 1858: 522. Leptodeira septentrionalis Kennicott, 1859: Figure 13A Material examined. Table 2. 226 Check List 13 (4)

Figure 12. Amphibian species present in Jaguas and San Carlos hydroelectric protected areas. A: Tantilla melanocephala MHUA-R 14855. B: Atractus occipitoalbus MHUA-R 15131. C: Clelia clelia. D: Coniophanes fissidens MHUA-R 14879. E: Erythrolamprus epinephelus MHUA-R 14841. F. Erythrolamprus melanotus. G: Erythrolamprus pseudocorallus MHUA-R 14856. H: Imantodes cenchoa MHUA-R 15134. Restrepo et al. | Herpetofauna of the northern Andes of Colombia 227

Figure 13. Amphibian species present in Jaguas and San Carlos hydroelectric protected areas. A: Leptodeira septentrionalis. B: Ninia atrata MHUA-R 14847. C: Ninia teresitae. MHUA-R 14860. D: Oxyrhopus petolarius MHUA-R 14843. E: Rhadinaea decorata MHUA-R 14859. F: Sibon nebulatus MHUA-R 15154. G: Siphlophis compressus MHUA-R 14733. H: Urotheca fulviceps MHUA-R 15132. 228 Check List 13 (4)

Micrurus mipartitus (Duméril, Bibron & Duméril, 1854): Pristimantis fallax, Pristimantis jaguensis, Pristimantis Figure 14C viejas, Rheobates pseudopalmatus, Rulyrana susatamai, Elaps mipartitus Duméril et al. 1854: 1220. and Sachatamia punctulata. The distribution of the Micrurus mipartitus — Amaral 1926: 66. other endemic species extends to the Magdalena river Material examined. Table 2. valley (Microcaecilia pricei, Caecilia thompsoni and Trilepida macrolepis (Peters, 1857): Figure 14D Craugastor metriosistus), the Chocoan Pacific lowland Stenostoma macrolepis Peters 1857: 402. (Colostethus inguinalis; Grant and Lynch 2004) or the Trilepida macrolepis— Hedges 2011: 63 [by implication]. Caribbean lowland and foothills of the Sierra Nevada Material examined. Table 2. de Santa Marta (Dendrobates truncatus). Among the This is a very small species that may be differentiated reptiles, Anolis sulcifrons was the only recorded spe- from other snakes by having body scales of the same size cies with a distribution restricted to Colombia (Uetz and (without enlarged ventral), arranged in 14 rows around Hošek 2016). de body (Köhler 2008). It has 4 supralabial scales and 10 Five records are of particular interest in this study. scale rows around the middle of the tail. Individuals are Pristimantis jaguensis is an endemic frog from Colom- dark brown dorsally and lighter ventrally, with whitish bia, with only 1 known locality, the surroundings of the scale borders (Pinto et al. 2010). Epictia goudotii, a spe- Jaguas hydroelectric project (Rivera-Prieto et al. 2014). cies potentially sympatric with Trilepida macrolepis, has Hyalinobatrachium aureogutattum is a frog previously 2 supralabials, a darker color pattern with stripes, and a recorded from the departments of Valle del Cauca, yellow rostral scale (Pinto et al. 2010). Risaralda, Chocó and Antioquia on the western flank of the Cordillera Occidental in Colombia, between 45 and Bothrops asper (Garman, 1883): Figure 14E 1570 m above sea level from the provinces of Esmer- Trigonocephalus asper Garman 1883: 124. Bothrops asper — Hoge 1966: 113. alda and Imbabura in Ecuador, and from the eastern Material examined. Table 2. cordilleras in southwestern Darién province in Panama (IUCN 2016). This is the first record of this species in Porthidium lansbergii (Schlegel, 1841): Figure 14F the Magdalena river valley in Colombia. The presence Trigonocephalus lansbergi Schlegel 1841: 1. of Rhadinaea decorata in Colombia was suggested by Porthidium lansbergii — McDiarmid et al. 1999: 320. Myers (1974), and we present a voucher of this spe- Material examined. Table 2. cies from the San Carlos site, which thus represents the Kinosternon leucostomum (Duméril, Bibron & Duméril, first verified record from Colombia. Anadia ocellata is 1851): Figure 14G distributed on both slopes of the cordilleras in Costa Cinosternon leucostomum Duméril et al. 1851: 17. Rica and western Panama (Savage 2002). Batista et al. Cinosternon postinguinale Cope 1887: 23. (2016) mention the occurrence in the Pacific lowlands Material examined. Table 2. of Colombia, contrary to Oftedal (1974), Savage (2002), This small freshwater cryptodiran turtle is yellowish and Köhler (2008), who indicated a southern range limit brown. It may be recognized by having a single carapace in Panama. We conservatively assign our voucher speci- keel and the axillar and inguinal scutes usually separated, men from the Magdalena river valley to the A. ocellata whereas Kinosternon scorpioides has 3 carapace keels group (sensu Oftedal 1974). The distribution of Rhinella and the axillar and inguinal scutes usually in contact alata in eastern cordillera was suggested by Pramuck (Köhler 2008). (2006), although her observation lacked a voucher. Here, we present vouchers of R. alata that confirm the inter- Discussion Andean distribution of this species with records from Magdalena river valley. The 43 amphibian species recorded in this study repre- Historically, the herpetofauna from the eastern side sent 5.2% of the Colombian amphibian species (Frost of the Andes has been considered very distinct from the 2016), and 15% of the species reported from the Cordil- trans-Andean region (Duellman 1979). Nevertheless, as lera Central in Colombia (IUCN 2016). The 58 reptile part of the long-term sampling carried out in this study, species represent 6% of the total Colombian reptiles we have recently reported on the occurrence of species species (Uetz and Hošek 2016), and 44% of the reptile distributed in the eastern flank of the northern Cordil- fauna reported for the Cordillera Central. The Cordillera lera Central but previously known from the Amazon Central in Colombia is recognized for its high diversity basin: Atractus occipitoalbus (Marin et al. 2017), Ano- in amphibians and reptiles (Lynch et al. 1997) and the 2 lis fuscoauratus (Grisales-Martínez et al. in press) and sites we studied represent a high proportion of this her- Cochranella resplendens (Molina-Zuluaga et al. 2017). petofauna. These findings urge the need for thorough sampling and Fifteen (35%) of the amphibian species found dur- a detailed taxonomic revision of species distributed in the ing the monitoring are endemic to Colombia. Most of Magdalena river valley to elucidate the biogeographic them are restricted to the eastern flank of the Cordil- connection between both areas. lera Central: Bolitoglossa ramosi, Dendropsophus We were not able to assign 7 records to any valid norandinus, Diasporus anthrax, Epicrionops parkeri, species. The record of Marisora sp. (Figure 9F) may Restrepo et al. | Herpetofauna of the northern Andes of Colombia 229

Figure 14. Amphibian species present in Jaguas and San Carlos hydroelectric protected areas. A: Xenodon rabdocephalus. B: Micru- rus dumerilii MHUA-R 15138. C: Micrurus mipartitus MHUA-R 15152. D: Trilepida macrolepis MHUA-R 14858. E: Bothrops asper. F: Porthi- dium lansbergii MHUA-R 14956. G: Kinosternon leucostomum. correspond to Marisora (Mabuya) candidate species IV specimen photograph (Figure 10B) we cannot assign it suggested by Pinto-Sánchez et al. (2015). The second to any of these species. The vouchers of Ptychoglossus record refers to an Ameiva sp. In Colombia, there are sp., Sachatamia sp., Colostethus aff. fraterdanieli, and 3 Ameiva species: Ameiva ameiva, A. bifrontata, and Cercosaura sp. may represent undescribed taxa. A. praesignis (Harvey et al. 2012). As we only have a Pristimantis fallax is the only species reported in this 230 Check List 13 (4)

Table 3. Taxonomic list of amphibians reported in the Jaguas and San Carlos Hydroelectric protected areas. Asterisks indicate species endemic to Colombia. IUCN Conservation status: CR = Critically Endangered, EN = Endangered, VU = Vulnerable, NT = Near Threatened, LC = Least Concern, DD = Data Deficient; NE = Not Evaluated. Geographic distribution follows Lynch et al. (1997): A = Chocoan lowland forests, B = Caribbean lowlands and inter-Andean valleys, C = Llanos, D and E = Amazonia, F = Cordillera Occidental, G = Cordillera Central, H = Cordillera Oriental, I = Sierra Nevada de Santa Marta.

Taxon Jaguas San Carlos IUCN Distribution AMPHIBIA ANURA Aromobatidae *Rheobates pseudopalmatus (Rivero & Serna, 2000) X X DD G Bufonidae Rhinella alata (Thominot, 1884) X X DD A Rhinella horribilis (Wiegmann, 1833) X X NE A-B, F-I Centrolenidae Cochranella resplendens (Lynch & Duellman, 1973) X LC G, D-E, Espadarana prosoblepon (Boettger, 1892) X X LC A–B, F-H Hyalinobatrachium aureoguttatum (Barrera-Rodríguez & Ruíz-Carranza, 1989) X NT A–B, G Hyalinobatrachium fleischmanni (Boettger, 1893) X LC A–B, F-H *Rulyrana susatamai (Ruiz-Carranza & Lynch, 1995) X VU B, G *Sachatamia punctulata (Ruiz-Carranza & Lynch, 1995) X X EN B, G Sachatamia sp. X Craugastoridae *Craugastor metriosistus Ospina-Sarria, Angarita-Sierra & Pedroza-Banda, 2015 X X NE B *Pristimantis fallax (Lynch & Rueda, 1999) X VU G Pristimantis gaigei (Dunn, 1931) X X LC A–B, F-H *Pristimantis jaguensis Rivera-Prieto , Rivera-Correa & Daza, 2014 X NE G Pristimantis penelopus (Lynch & Rueda, 1999) X X VU B, G–H Pristimantis taeniatus (Boulenger, 1912) X X LC A–B, F–H *Pristimantis viejas (Lynch & Rueda-Almonacid, 1999) X X LC B, G–H Dendrobatidae *Colostethus inguinalis (Cope, 1868) X LC A–B, G Colostethus aff. fraterdanieli X *Dendrobates truncatus (Cope, 1861) X X LC A–B, G–I Eleutherodactylidae *Diasporus anthrax (Lynch, 2001) X X DD B, G Diasporus gularis (Boulenger, 1898) X X LC A–B, F–G Hemiphractidae Hemiphractus fasciatus Peters, 1862 X NT A–B, F Hylidae Boana boans (Linnaeus, 1758) X X LC A–I Boana xerophylla (Duméril & Bibron, 1841) X X NE B–C, F–I Dendropsophus ebraccatus (Cope, 1874) X X LC A–B, G–H Dendropsophus microcephalus (Cope, 1886) X X LC A–C, G–I *Dendropsophus norandinus Rivera-Correa & Gutiérrez-Cárdenas, 2012 X NE G Dendropsophus subocularis (Dunn, 1934) X X LC A–B, F–H Hyloscirtus palmeri (Boulenger, 1908) X X LC A–B, F–H Phyllomedusa venusta Duellman & Trueb, 1967 X X LC B, G–I Scinax sp. X X Scinax rostratus (Peters, 1863) X X LC A–C, F–I Scinax ruber (Laurenti, 1768) X X LC A–I Smilisca phaeota (Cope, 1862) X X LC A–B, F–G Leptodactylidae Engystomops pustulosus (Cope, 1864) X X LC A–C, G–I Leptodactylus colombiensis Heyer, 1994 X X LC B–C, F–H Leptodactylus fuscus (Schneider, 1799) X X LC B–I Leptodactylus savagei Heyer, 2005 X LC B,F-G, I CAUDATA Plethodontidae *Bolitoglossa ramosi (Brame & Wake, 1972) X LC G GYMNOPHIONA Caeciliidae *Caecilia thompsoni (Boulenger, 1902) X X DD B, G–H Siphonopidae *Microcaecilia pricei X LC B,F Rhinatrematidae *Epicrionops parkeri (Dunn, 1942) X DD G Restrepo et al. | Herpetofauna of the northern Andes of Colombia 231

Table 4. Taxonomic list of reptiles reported in the Jaguas and San Carlos Hydroelectric protected areas. Asterisks indicate species endemic to Colombia. IUCN Conservation status is shown for both: a global assessment and a national assessment (IUCN 2016/Morales-Betancourt et al. 2015). CR= Critically Endangered, EN = Endangered, VU = Vulnerable, NT = Near Threatened, LC = Least Concern, DD = Data Deficient; NE = Not Evaluated. Geographic distribution follows Lynch et al. (1997): A = Chocoan lowland forests, B = Caribbean lowlands and inter-Andean valleys, C = Llanos, D and E = Amazonia, F = Cordillera Occidental, G = Cordillera Central, H = Cordillera Oriental, I = Sierra Nevada de Santa Marta.

IUCN Taxon Jaguas San Carlos (Global/ Distribution National) REPTILIA SQUAMATA SAURIA Corytophanidae Basiliscus galeritus Dúmeril, 1851 X X NE/LC A–B, G Corytophanes cristatus (Merrem, 1820) X X LC/LC A–B, G Dactyloidae Anolis auratus Daudin, 1802 X NE/LC A–H Anolis aff. granuliceps X X Anolis fuscoauratus D’Orbigny, 1837 X NE/LC *Anolis sulcifrons Cope, 1899 X X NE/NT B, G–H Anolis tropidogaster Hallowell, 1856 X NE/LC Anolis vittigerus Cope 1862 X NE/LC A-B, G–H Gymnophthalmidae Anadia gr. ocellata Gray, 1845 X NE/NT A–B, G Cercosaura sp. X X Echinosaura palmeri Boulenger, 1911 X X LC/NE A, G Loxopholis rugiceps Cope, 1869 X LC/LC A-B, G-I Loxopholis southi Ruthven & Gaige, 1924 X LC/LC A, G Ptychoglossus sp. X X Phyllodactylidae Thecadactylus rapicauda (Houttuyn, 1782) X X NE/LC A–C, E, G–H Polychrotidae Polychrus gutturosus Berthold, 1846 X X NE/LC A–B, F–H Scincidae Marisora sp. X X Sphaerodactylidae Gonatodes albogularis (Duméril & Bibron, 1836) X X NE/LC A–I Lepidoblepharis xanthostigma (Noble, 1916) X X LC/LC A–B, G Sphaerodactylus lineolatus Lichtenstein & Von, 1856 X LC/LC A–B, G Teiidae Ameiva sp. X X Cnemidophorus lemniscatus (Linnaeus, 1758) X X NE/NE A–B, D–E, G Holcosus festivus (Lichtenstein, 1856) X X LC/LC A–B, G–H AMPHISBAENIA Amphisbaenidae Amphisbaena varia Laurenti, 1768 X X NE/LC A–H OPHIDIA Boidae Boa constrictor Linnaeus, 1758 X X NE/LC A–C, E, G–H Colubridae Chironius exoletus (Linnaeus, 1758) X X NE/LC A–H Chironius grandisquamis (Peters, 1869) X X LC/LC A–B, F–G Dendrophidion percarinatum (Cope, 1893) X X LC/LC A–B, F–G Drymobius rhombifer (Günther, 1860) X LC/LC A–I Leptophis ahaetulla (Linnaeus, 1758) X X NE/LC A–I Mastigodryas boddaerti (Sentzen, 1796) X X NE/LC A–I Oxybelis aeneus (Wagler, 1824) X X NE/LC A–H Phrynonax poecilonotus (Günter, 1858) X X LC/NE A–B, F–H Scaphiodontophis venustissimus (Gunther, 1893) X LC/NE G Spilotes pullatus (Linnaeus, 1758) X X NE/LC A–I Tantilla melanocephala (Linnaeus, 1758) X NE/LC A–C, E–H Dipsadidae Atractus occipitoalbus (Jan, 1862) X NT/NT E, G Clelia clelia (Daudin, 1803) X X NE/LC A–I Coniophanes fissidens (Gunther, 1858) X X NE/LC B, F–G Erythrolamprus epinephelus (Cope, 1862) X NE/LC F–I Erythrolamprus melanotus (Shaw, 1802) X LC/LC A–I Erythrolamprus pseudocorallus Roze, 1959 X X LC/LC B, G–H Imantodes cenchoa Linnaeus, 1758 X X NE/LC A–H Continued 232 Check List 13 (4)

Table 4. Continued.

IUCN Taxon Jaguas San Carlos (Global/ Distribution National) Leptodeira septentrionalis Kennicott, 1859 X X NE/LC A–B, F–H Ninia atrata (Hallowell, 1845) X X LC/LC B–C, E–H *Ninia teresitae Angarita-Sierra & Lynch, 2017 X NE/NE B Oxyrhopus petolarius (Linnaeus, 1758) X X NE/LC A–H Rhadinaea decorata (Gunther, 1858) X X NE/NE G Sibon nebulatus (Linnaeus, 1758) X X NE/LC A–C, F–I Siphlophis compressus (Daudin, 1803) X LC/LC A–E, G Urotheca fulviceps (Cope, 1886) X NE/DD A–B, H Xenodon rabdocephalus (Wied-Neuwied, 1824) X NE/LC A–B, E–H Elapidae Micrurus dumerilii Jan, 1858 X NE/LC A–B, F–I Micrurus mipartitus (Duméril, Bribon & Duméril, 1854) X NE/LC A–B, F–I Leptotyphlopidae Trilepida macrolepis (Peters, 1857) X NE/LC A–B, D–E, G–H Viperidae Bothrops asper (Garman, 1883) X X NE/LC A–B, G–H Porthidium lansbergii (Schlegel, 1841) X X NE/LC A–B, F–H TESTUDINES Kinosternidae Kinosternon leucostomum (Duméril, Bibron & Duméril, 1851) X X NE/LC A–B, G–H study included as Vulnerable at the country level in the ity will give us a better picture of the regional diversity Resolucion 192 del 2014 del Ministerio de Ambiente y and will impact future studies aiming to understand the Desarrollo Sostenible. At the global level, Sachatamia origin and maintenance of this rich fauna. punctulata is listed by the IUCN as Endangered because The high species density, high representativeness of its extent of occurrence is less than 5000 km2, its distri- the regional biodiversity, and the high levels of amphib- bution is severely fragmented, and there is a continuing ian endemism make these 2 private protected areas very decline in the extent and quality of its habitat (IUCN important for conservation of the regional herpetofauna. 2016). Pristimantis fallax, P. penelopus and Rulyrana As shown here, compensation areas of large infrastructure susatamai are listed as Vulnerable because of continuous projects (i.e., hydroelectric projects) represent a potential decline in the extent and quality of their habitat (Castro et opportunity to preserve the already highly degraded eco- al. 2004, IUCN 2016). Lastly, 2 species are categorized systems in the northern Andes of Colombia. as Near Threatened: Hemiphractus fasciatus Peters, 1862 and Hyalinobatrachium aureoguttatum (Coloma et al. Acknowledgements 2008, Solís et al. 2010; Table 3). Recently, the conservation status of Colombia’s rep- During almost a decade, ISAGEN and the Universidad tiles has been assessed and 43 species are now under de Antioquia have funded herpetofaunal studies in this one of the 3 threat categories (Morales-Betancourt et region. This manuscript was partially funded under Isa- al. 2015). However, none of them correspond to species gen project 47/574. We thank the Grupo Herpetológico present in our study areas. It is well known that reptiles de Antioquia and all the herpetologists and students who around the world are declining due to a number of causes over the years have been collecting information about directly related to human activities (Gibbons et al. 2000). the amphibian and reptile diversity in this highly diverse In addition to habitat loss and degradation, another threat region of the Andes. M. Rivera-Correa provided useful is present in Colombia: the cultural view that reptiles, in comments to the manuscript. particular snakes, are dangerous and nasty animals that should be exterminated (Castaño-Mora 2002, Lynch Authors’ Contributions 2012). This view has probably driven several snake species to the brink of extinction at least locally (Lynch AR, CMZ and JMD conceived the idea and wrote the 2012). text. JMD, CMM and JPH confirmed the taxonomic To build complete species lists for Colombian identity of all specimens. amphibians and reptiles evokes large challenges for the following reasons: (1) the high levels of species diversity References of amphibians and reptiles in the area, (2) the extreme cryptic diversity in several genera (e.g., Pristimantis, Acevedo AA, Lampo M, Cipriani R (2016) The cane or marine toad, Rhinella marina (Anura, Bufonidae): two genetically and morpho- Anolis, Atractus), and (3) the lack of well-identified refer- logically distinct species. Zootaxa 4103 (6): 574–586. http://doi. ence collections. However, continued specimen collection org/10.11646/zootaxa.4103.6.7 throughout different seasons and years in a specific local- Angarita-Sierra T, Lynch JD (2017) A new species of Ninia (Serpen- Restrepo et al. | Herpetofauna of the northern Andes of Colombia 233

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